Article

Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae)

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Abstract

Minuartia is one of the larger genera of Caryophyllaceae with about 175 species distributed mainly in the Northern Hemisphere. The taxonomy of the genus has been studied by several authors, resulting in the recognition of numerous infrageneric groups. Previous phylogenies of the Caryophyllaceae showed that Minuartia is polyphyletic, but included only a limited sample of the genus. We here provide a molecular phylogeny of Minuartia including all infrageneric groups recognized in the last revision of the genus. We reconstructed the phylogeny of the genus using DNA sequences of nrITS and plastid matK from 160 ingroup and 105 outgroup samples. The evolution of the morphological character relevant for the delimitation of Minuartia (three styles plus three capsule valves/teeth) was reconstructed. Minuartiahas been defined with a plesiomorphic character and is highly polyphyletic. All four subgenera fall into different lineages containing other genera of the family, and M. subg. Minuartia, as by far the largest subgenus, falls into seven clades, which together do not form a monophylum. These clades are closely related to several other genera, e.g., Sagina, Colobanthus and Scleranthus. In several cases taxonomic groups below subgeneric rank are monophyletic. The type of Minuartia, M. dichotoma, is part of a clade containing M. sect. Plurinerviae and sect. Minuartia. We propose to retain this clade as Minuartia s.str. and to transfer the other species of Minuartia to the genera Cherleria, Eremogone, Facchinia, Mcneillia, Minuartiella, Mononeuria, Pseudocherleria, Rhodalsine, Sabulina and Triplateia.

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... However, this was not due to a lack of discriminatory power in the DNA regions used in this study. Molecular phylogenies have previously suggested that these two genera are polyphyletic [44,61]. In the case of Minuartia, the three species (Minuartia arctica, Minuartia laricina, and Minuartia verna (L.) Hiern var. ...
... leptophylla (Rchb.) Nakai) analyzed in this study each belong to different sections within the genus (summarized in [61]). The molecular phylogeny also showed that each of these species are included in different clades [61]. ...
... Nakai) analyzed in this study each belong to different sections within the genus (summarized in [61]). The molecular phylogeny also showed that each of these species are included in different clades [61]. In fact, Minuartia laricina and Minuartia verna var. ...
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Caryophyllaceae is a large angiosperm family, with many species being utilized as ornamental or medicinal plants in Korea, in addition to several endangered species that are managed by the government. In this study, we used DNA barcoding for the accurate identification of Korean Caryophyllaceae. A total of 78 taxa (n = 215) were sequenced based on three chloroplast regions (rbcL, matK, and psbA–trnH) and nuclear ribosomal internal transcribed spacers (ITS). In the neighbor-joining tree, a higher accuracy of identification was generally observed when using ITS (>73%) rather than chloroplast regions (<62%). The highest resolution was found for rbcL + ITS (77.6%), although resolution varied according to the genus. Among the genera that included two and more species, five genera (Eremogone, Minuartia, Pseudostellaria, Sagina, and Stellaria) were successfully identified. However, the species of five other genera (Cerastium, Gypsophila, Dianthus, Silene, and Spergularia) showed relatively low resolutions (0–61.1%). In the cases of Cerastium, Dianthus, and Silene, ambiguous taxonomic relationships among unidentified species may have been a factor contributing to such low resolutions. However, in contrast to these results, Gypsophila and Spergularia have been identified well in previous studies. Our findings indicate the need of taxonomic reconsideration in Korea.
... Two species are growing in South America. It is highly variable in chromosome number and morphologically defined by linear to linear-setaceous leaves, pentamerous flowers, acute and usually three-veined sepals, a calyx that is not hardened at the base, white petals, three styles, and three capsule valves (Mattfeld, 1922;Dillenberger & Kadereit, 2014;Dillenberger, 2016). ...
... The genus Sabulina was first described by Reichenbach (1832) but was later included in Minuartia as section Sabulina by Ascherson & Graebner (1918). Recent molecular approaches have shown that endemism rate of 61% (Koç, 2012;Dillenberger & Kadereit, 2014;Koç et al., 2019Koç et al., , 2021. ...
... The voucher numbers and locality information of the specimens investigated were given in Table 1. The order of the taxa was adopted from Dillenberger & Kadereit (2014) and Koç et al. (2019Koç et al. ( , 2021. ...
Article
Pollen morphology of 28 taxa belonging to the genus Sabulina Rchb. (Caryophyllaceae) from Turkey was examined and compared using light microscopy and scanning electron microscopy. The current study aimed to describe the palynological characteristics of the Sabulina taxa and to assess the diagnostic value of these features for taxonomic implications. The pollen grains of Sabulina are radially symmetrical, apolar, rarely isopolar, and pantopolyporate with 8–28 pores. Their shape is prolate-spheroidal with the A axes (long axes) 19.5 μm and the B axes (short axes) 19–39.5 μm. Exine sculpturing is microechinate-punctate in all taxa. Principal component analysis was used to evaluate the significant pollen morphological characters for distinguishing the taxa, with three components accounting for 95.112% of the total variance. Numerical analysis revealed that pollen sizes, pore diameters, pore numbers, distance between pores, spinule density, and punctum numbers are particularly the most useful variables for separating the Sabulina taxa. The results also display a partial relationship between the Sabulina taxa clusters based on pollen morphology and the systematics of these taxa based on general macromorphology.
... s.l. [26]. ...
... Dillenberger and Kadereit [26] documented rampant polyphyly in Minuartia as traditionally conceived, revealing that the main diagnostic characters for the genus (i.e., presence of three styles and three fruit valves) are indeed plesiomorphic. Even subgeneric ranks (e.g., Minuartia subg. ...
... A synopsis of the most relevant taxonomic treatments is provided in Table 1. In addition, the relationships between the taxa are still largely speculative (e.g., [27]), and the phylogeny almost unknown; as at present only four species (mostly with single specimens) have been investigated by molecular methods [26]. In this study, we infer nuclear and plastid phylogenies of genus Mcneillia involving all its species and subspecies across their geographic distributions. ...
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The genus Mcneillia has been recently segregated from Minuartia L. based on molecular results, also supported by morphology. However, to date, a comprehensive study on the phylogenetic relationships within this genus is lacking. In this paper, we provide a multigene phylogeny of all the species and subspecies of Mcneillia employing two nuclear and six chloroplast markers. We documented extensive gene flow between taxa, sometimes separated at specific rank. In addition, Mcneillia as currently circumscribed, is not monophyletic. In fact, Mcneillia graminifolia subsp. brachypetala, strictly endemic to Greece, truly belongs to Minuartiella, a genus otherwise limited to South-West Asia. Moreover, even after removal of this taxon, our results do not support the monophyly of the taxa included in M. graminifolia s.l., the most variable and widespread species of the genus. Further controversial subspecies of Mcneillia graminifolia, i.e., subsp. hungarica and subsp. rosanoi, are shown to deserve taxonomic recognition as separate species, whereas Mc. moraldoi is not distinct at specific rank. In addition, Mc. saxifraga subsp. tmolea is here regarded as a further distinct species. A consistent taxonomic treatment is therefore proposed with six new combinations and nomenclatural notes, providing the necessary typifications.
... Arenaria (Sadeghian & al., 2015) and Minuartia Loefl. (Dillenberger & Kadereit, 2014). Sadeghian & al. (2015) confirmed the exclusion of the majority of Arenaria spp. ...
... Odontostemma to genus rank, maintaining them as members of Alsineae. Minuartia, also treated as a member of Alsineae in traditional systems (Pax & Hoffmann, 1934;Schischkin, 1936;Bittrich, 1993), turned out to be polyphyletic with members placed in both tribes Sagineae J.Presl and Sclerantheae DC. (Dillenberger & Kadereit, 2014). A recent study of Pseudostellaria retrieved this genus as a member of the Odontostemma clade, and the new genus Hartmaniella M.L.Zhang & Rabeler was proposed for two American species formerly assigned to Pseudostellaria (Zhang & al., 2017). ...
... -The placement of a few species of Stellaria, i.e., St. ovata (Mexico to S America), St. minutifolia, and St. howardii (Dominican Republic), among the outgroups is in agreement with previous molecular phylogenies (Harbaugh & al., 2010;Greenberg & Donoghue, 2011;Dillenberger & Kadereit, 2014). Morphological and geographic evidence (growth habit, 4-merous flowers, relatively shallowly cleft petals, seed with prominent dorsal tubercles, and distribution in Antillean islands) suggests St. antillana Urb. is closely allied to St. ovata (Maguire, 1958) and belongs to this group, which was also confirmed by a recent phylogenetic analysis (Sharples & Tripp, 2019a). ...
Article
The tribe Alsineae is a large monophyletic group in the family Caryophyllaceae especially found across Eurasia and the Americas, but with a center of diversity in the Mediterranean region. Several previous molecular phylogenetic studies have focused on the delimitation of genera and tribes of Caryophyllaceae, especially the subfamily Alsinoideae or the tribe Alsineae in a broader sense than now recognized. However, there are still many open questions regarding the subdivision of the tribe and genus delimitation. In the present study, we sampled 191 (148 species) and 149 (125 species) accessions of Alsineae representing almost all (Adenonema and Pseudocerastium were not available to us) recognized genera in the tribe for nuclear DNA internal transcribed spacer (ITS) and plastid marker rps16 sequences, respectively. A combined matrix of 103 species was built for taxa with both sequences available. Maximum parsimony and Bayesian inference analyses retrieved Cerastium and Stellaria (including Myosoton) as the largest monophyletic genera, while other genera were medium‐sized (10–20 spp.) or small (<10 spp.). Our expanded sampling of Pseudostellaria and its relatives suggests a broader circumscription of this genus. Major divergence in morphology, particularly of the seeds, observed in the “Protostellaria”‐clade, allows recognition of some taxonomic changes. A total of 16 genera are recognized in Alsineae including Cerastium, Dichodon, Hartmaniella, Holosteum, Lepyrodiclis, Mesostemma, Moenchia, Nubelaria, Odontostemma, Pseudostellaria, Rabelera, Schizotechium, Shivparvatia, and Stellaria, along with Adenonema and Pseudocerastium that could not be analyzed and are, therefore, kept as distinct genera. A diagnostic key to these genera, as well as notes on their relationships, distribution, and nomenclature, is provided.
... The genus Minuartia L. (Family: Caryophyllaceae) is the particularly of interest because of its preference for metalliferous and other extreme substrates. This genus, previously included in Arenaria (Maguire 1951), comprises > 170 species distributed mainly in the Northern Hemisphere, between temperate and arctic-alpine region (Dillenberger and Kadereit 2014). One of the main centers of Minuartia diversity is Europe (85 taxa), especially the Mediterranean region (Halliday 1993;Diklić and Stevanović 2012), with a significant number of species in the Balkan Peninsula (47 taxa; Diklić and Stevanović 2012). ...
... Graebn., generally calcifuge species from the monophyletic Sect. Plurinervie McNeill (Dillenberger and Kadereit 2014). Although also considered a subspecies (M. ...
... Stoj. & Stef.; Dillenberger and Kadereit 2014), M. bulgarica is an accepted taxon according to relevant databases (Euro+Med 2006; http:// www. thepl antli st. ...
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The aim of this study was to determine the concentrations of potentially toxic elements in soil samples and plant tissues of Minuartia recurva and M. bulgarica, predominantly or exclusively calcifuge species. Biological concentration (BCs) and translocation factors (TFs) were used to evaluate their accumulation potential. Considerable differences were observed between M. recurva and M. bulgarica assessions in terms of accumulation strategies of potentially toxic elements (PTEs). In M. recurva, most of the elements analyzed (Mn, Cu, Zn, Cd, and Co) were transported to the shoot, whereas in M. bulgarica, these elements remained predominantly in the roots. The Cu concentrations in the shoot samples of M. recurva from an abandoned iron-copper mine at Mt. Kopaonik were clearly above the notional hyperaccumulation threshold, characterizing this species as a possible Cu hyperaccumulator. Additionally, strong accumulation potential for Cr, Ni, Zn, Pb, and Cd was observed in M. recurva assessions, but without significant accumulation due to the low concentrations of these elements in the soils. The strong accumulation capacity and the different strategies in tolerance to PTEs indicate a potential of the two species for an application in phytoremediation: M. recurva for phytoextraction and M. bulgarica for phytostabilization.
... In Sagina, outer stamens and capsule valves are episepalous and inner stamens and styles are alternisepalous (Crow 1978;Timaná 2018). Although the most comprehensive phylogeny including Colobanthus and Sagina revealed the two genera to be monophyletic, as assumed by Bittrich (1993), it contained only four species of each genus (Dillenberger and Kadereit 2014). Although Pufal (2010) analyzed a larger species sample of Colobanthus, the sequences used are not publicly available. ...
... The closest relatives of Sagina and Colobanthus, i.e. Facchinia and Sabulina Rchb., (Dillenberger and Kadereit 2014) are predominantly Northern Hemisphere genera with together more than 80 species. Two other genera, Bufonia L., and Drypis L., possibly also belonging to tribe Sagineae sensu Harbaugh et al. (2010), are found only in the Northern Hemisphere. ...
... These DNA regions had proved to be phylogenetically informative in Caryophyllaceae in previous studies (e.g. Dillenberger and Kadereit 2014;Zhang et al. 2016;Legler and Dillenberger 2017;Biersma et al. 2020). Polymerase chain reactions (PCR) for ITS and trnQ-rps16 were carried out as described in Kadereit (2014, 2017) and Biersma et al. (2020). ...
Article
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Colobanthus (23 species) and Sagina (30–33 species) together are sister to Facchinia. Whereas Facchinia is distributed in western Eurasia, Colobanthus is almost exclusively distributed in the Southern Hemisphere, and Sagina is distributed in both hemispheres with the highest species diversity in western Eurasia. We examined: 1. Whether Sagina and Colobanthus are monophyletic sister genera, 2. Where the two genera originated and how many times dispersal between hemispheres occurred, and 3. Which colonization routes between hemispheres were taken. We reconstructed the phylogeny of Colobanthus and Sagina using nuclear ribosomal internal transcribed spacer (ITS) and two plastid spacers (cpDNA) of altogether 158 ingroup samples of 45 species, and performed molecular dating and ancestral area reconstructions. Sagina and Colobanthus were confirmed as monophyletic sister genera. Biogeographical reconstructions based on ITS and cpDNA showed that Sagina reached the Southern Hemisphere in Australasia or in Africa. For Colobanthus, patterns were less clear and less well-supported: ITS showed Australasia as the region of entry, but cpDNA implied that the Southern Hemisphere may have been entered in America. The extant distributions and the biogeographical histories of Colobanthus and Sagina show both similarities and dissimilarities. This illustrates that biogeographical histories, even of closely related and ecologically very similar lineages, can be highly idiosyncratic.
... Greenberg and Donoghue 2011). At genus rank, several studies have been carried out on Arenaria L., Minuartia L., Dianthus L., Gypsophila L., Polycarpon L., Silene L., etc. (see e.g., Kool et al. 2007, Iamonico 2013, 2015, 2016, Dillenberger and Kadereit 2014, Iamonico and Domina 2015, Sadeghian et al. 2015, Dillenberger and Rabeler 2018, Madhani et al. 2018), but various questions are still open. ...
... This clade corresponds to the tribe Sagineae. Dillenberger and Kadereit (2014), who investigated in detail the genus Minuartia, did not consider the genus Habrosia in their analysis. However, the species of Minuartia, included in the tribe Sagineae by Greenberg and Donoghue (2011), were treated by Dillenberger and Kadereit (2014) as belonging to the resurrected genus Sabulina Rchb. ...
... Dillenberger and Kadereit (2014), who investigated in detail the genus Minuartia, did not consider the genus Habrosia in their analysis. However, the species of Minuartia, included in the tribe Sagineae by Greenberg and Donoghue (2011), were treated by Dillenberger and Kadereit (2014) as belonging to the resurrected genus Sabulina Rchb. In contrast to Habrosia, Dillenberger and Kadereit (2014) included Arenaria fontinalis in their analysis, which was also investigated by Greenberg and Donoghue (2011), and confirmed that it is to be treated as a member of Sabulina and, in fact, a new combination, S. fontinalis (Short and R. Peter) Dillenb. ...
Article
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Habrosia (Sagineae, Caryophyllaceae) is a genus that includes only H. spinuliflora, a species occurring in Iran, Iraq, Syria, Lebanon, and Turkey (Irano-Turanian floristic chorological element). Based on the available molecular data published in 2011, Habrosia appears to be nested in a Minuartia-clade, which includes taxa currently recognized under the genus Sabulina. Consequently, Habrosia should be treated as a genus to be included in Sabulina. However, the molecular tree published in 2011 considered only 9 Sabulina members whereas, according to the current concept, Sabulina is a genus comprising about 65 species. Unfortunately, the molecular phylogeny including a larger Sabulina sample published in 2014 did not include H. spinuliflora and the taxonomic position of Habrosia remains, therefore, uncertain. With the aim of verifying the correct position of Habrosia in the tribe Sagineae with respect to its relationship to Sabulina, a comprehensive molecular investigation based on ITS sequences, linked to detailed morphological data, is presented. The results obtained revealed that Habrosia is not part of Sabulina. A detailed description of H. spinuliflora, its ecological preference, and a distribution map are provided. Eventually, the name Arenaria spinulifolia (basionym of H. spinuliflora) is lectotypified on a specimen preserved at G (barcode G00212963).
... while Gucel (2013) recorded 2n=30 in M. nifensis. Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). ...
... Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). Ghaffari & Kelich (2006) studied M. lineate and reported a value of n=10. ...
... Our cytological studies are based on six taxa comprising Minuartia. M. picta are showing the dibasic nature of the genus with x=11 and 14, this result agreed with the findings of Dillenberger & Kadereit (2013) who recorded a value of x=11, and our In this study, the chromosome numbers, karyotypes, ideograms, and karyotype asymmetry degrees of Minuartia were determined. ...
Article
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Chromosome count, karyotypic character analysis, meiotic studies, monoploid karyograms and ideograms were performed in six taxa of Minuartia growing in Iraq (M. hamata, M. hybrida subsp. hybrida, M. intermedia, M. meyeri, M. picta and M. hybrida subsp. turcica). Species of M. hamata and M. meyeri showed 2n=2x=30 chromosome number, while M. hybrida subsp. hybrida and M. intermedia were diploid (26). The chromosome number (n=x) of six species was studied, and was found to be n=15 in M. hamata and M. meyeri, 13 in M. hybrida and M. intermedia, while in M. picta we recorded values of n= 11 and 14. Karyotype analysis of this species was first carried out in our study. Analysis of metaphases showed that the karyotype formula was mainly metacentric, submetacentric, and sub acrocentric. The sizes of the chromosomes were mainly small and very small. The course of meiosis varied from normal to abnormal. Abnormal microsporogenesis formation of two bridge chromosomes was detected in M. hamata and one bridge chromosome in M. intermedia and M. meyeri. Formation of laggard’s chromosomes was detected in M. hamata, M. meyeri and M. intermedia. As well as ring chromosome was showed in M. hybrida subsp. hybrida, also, some cells contain triad cell in metaphase stage instead four cells, as well as founded cell, contains two nuclei in same species which led to reduced pollen fertility and differences in pollen grain size.
... nemzetség a Caryophyllaceae Juss. család Alsinoideae Fenzl alcsaládjába tartozik (BITTRICH 1993), mintegy 175 fajt tartalmaz (DILLENBERGER & KADEREIT 2014). A nemzetség fajai főként az északi félteke mérsékelt égövi régióiban honosak (MCNEILL 1962) Minuartia tenuifolia (L.) Hiern, Minuartia hybrida (Vill.) ...
... néven tárgyalta a taxont (pl. PIGNATTI 1982, FAVARGER & MONTSERRAT 1990, HALLIDAY 1993, JAU-ZEIN 1995, STRID & TAN 1997, RUTKOWSKI 2008, LAMBION & VERLOOVE 2012, SÂRBU et al. 2013, TISON & DE FOUCAULT 2014, azonban a legújabb nevezéktani és genetikai vizsgálatok (DILLENBERGER & KADEREIT 2014, IAMONICO 2014, DILLENBERGER 2016) nyomán a faj jelenleg elfogadott neve Sabulina tenuifolia (L.) Rchb. (POWO 2024). ...
Article
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Railways are important introduction and dispersal corridors for plant species. The first establishment and spread of many alien plant species in Hungary can be linked to railways nowadays. Sabulina tenuifolia (L.) Rchb. is a sub-Atlantic–sub-Mediterranean–Western-Asian plant, which was not previously known from the flora of Hungary, but a small population of the species was found in Budapest, at the Rákospalota-Újpest railway station. The plant was probably introduced as a stowaway of rail transport and its further spread along railways is expected. The inclusion of the species in the Hungarian plant identification key has been published in the article.
... When used in electronics, these kinds of materials offer numerous benefits and allow for quick design and manufacturing procedures. For instance, DNA's use in semiconductor devices like diodes and solar devices have been thoroughly researched [23]. ...
... Fior et al. (2006) reported that subfamilies are polyphyletic in their study based on matK and ITS data [18]. With the conducted studies, taxa previously classified under different genera, including the Arenaria subgenus Eremogone (approximately 70 species), Arenaria subgenus Eremogoneastrum Williams (22 species), and Minuartia subgenus Spergella (Fenzl) McNeill (3 species), have been transferred to the genus Eremogone [19][20][21][22][23][24]. ...
... Arenaria L. (Caryophyllaceae Juss.) is a genus comprising about 160-175 species of annual and perennial herbs mostly distributed in the northern temperate and subarctic regions, the Mediterranean, Mexico, and the Andes of South America (see Bittrich 1993;Hernández-Ledesma et al. 2015;POWO 2023) but other authors (Sadeghian et al. 2015) include 150-300 species in the genus. The molecular data by Greenberg and Donoghue (2011), Dillenberger and Kadereit (2014), and Sadeghian et al. (2015) showed that Arenaria is polyphyletic and several names were transferred to other genera (see e.g., Conti et al. 2014;Dillenberger and Kadereit 2014;Sadeghian et al. 2015;Iamonico 2016;De Luca et al. 2022). From a nomenclatural point of view, questions remaining to be addressed concern several names that are still untypified (see e.g., Iamonico 2013Iamonico , 2014Iamonico , 2016Iamonico , 2019Iamonico , 2022. ...
... Arenaria L. (Caryophyllaceae Juss.) is a genus comprising about 160-175 species of annual and perennial herbs mostly distributed in the northern temperate and subarctic regions, the Mediterranean, Mexico, and the Andes of South America (see Bittrich 1993;Hernández-Ledesma et al. 2015;POWO 2023) but other authors (Sadeghian et al. 2015) include 150-300 species in the genus. The molecular data by Greenberg and Donoghue (2011), Dillenberger and Kadereit (2014), and Sadeghian et al. (2015) showed that Arenaria is polyphyletic and several names were transferred to other genera (see e.g., Conti et al. 2014;Dillenberger and Kadereit 2014;Sadeghian et al. 2015;Iamonico 2016;De Luca et al. 2022). From a nomenclatural point of view, questions remaining to be addressed concern several names that are still untypified (see e.g., Iamonico 2013Iamonico , 2014Iamonico , 2016Iamonico , 2019Iamonico , 2022. ...
Article
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The names Arenaria mattfeldii , A. pallens , A. peruviana , A. pintaudii , and A. stuebelii (Caryophyllaceae, Arenarieae) from Peru and Bolivia were studied and neotypified based on specimens preserved at B and P.
... In the past two decades, molecular phylogenetic studies have changed our understanding on the circumscription of the tribe Alsineae as well as generic delimitation within it (Fior et al., 2006;Harbaugh et al., 2010;Greenberg and Donoghue, 2011;Dillenberger and Kadereit, 2014;Pusalkar and Singh, 2015;Sadeghian et al., 2015;Zhang et al., 2017;Sharples and Tripp, 2019;Yao et al., 2021;Arabi et al., 2022;Wang et al., 2023). Bittrich (1993) provided a comprehensive taxonomic treatment of Alsineae and accepted 23 genera in the tribe, viz. ...
... For instance, Honkenya, Schiedea (including Alsinidendron; Weller et al., 1995;Hernańdez-Ledesma et al., 2015) and Wilhelmsia were transferred to the tribe Sclerantheae (Harbaugh et al., 2010;Greenberg and Donoghue, 2011); Bufonia, Colobanthus and Sagina were moved to Sagineae (Harbaugh et al., 2010;Greenberg and Donoghue, 2011); the genus Moehringia and several subgenera of Arenaria sensu lato (s.l.) were placed in Arenarieae (Harbaugh et al., 2010;Greenberg and Donoghue, 2011); and the genus Thylacospermum was placed in Thylacospermeae (Pusalkar, 2015;Xu et al., 2019). In addition, species of Minuartia have been segregated into 11 genera, all of which have been removed from Alsineae (Dillenberger and Kadereit, 2014). ...
Article
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Alsineae are one of the most taxonomically difficult tribes in Caryophyllaceae and consist of over 500 species distributed in the northern temperate zone. Recent phylogenetic results have improved our understanding on the evolutionary relationships among Alsineae members. Nevertheless, there are still some unresolved taxonomic and phylogenetic problems at the generic level, and the evolutionary history of major clades within the tribe was unexplored to date. In this study, we carried out phylogenetic analyses and divergence time estimation of Alsineae using the nuclear ribosomal internal transcribed spacer (nrITS) and four plastid regions (matK, rbcL, rps16, trnL-F). The present analyses yielded a robustly supported phylogenetic hypothesis of the tribe. Our results showed that the monophyletic Alsineae are strongly supported to be the sister of Arenarieae, and the inter-generic relationships within Alsineae were mostly resolved with strong support. Both molecular phylogenetic and morphological evidence supported the Asian species Stellaria bistylata and the two North American species Pseudostellaria jamesiana and Stellaria americana all should be recognized as new monotypic genera respectively, and three new genera Reniostellaria, Torreyostellaria, and Hesperostellaria were thereby proposed here. Additionally, molecular and morphological evidence also supported the proposal of the new combination Schizotechium delavayi. Nineteen genera were accepted within Alsineae and a key to these genera was provided. Molecular dating analysis suggested that Alsineae splitted from its sister tribe at ca. 50.2 million-years ago (Ma) during the early Eocene and began to diverge at ca. 37.9 Ma during the late Eocene, and divergent events within Alsineae occurred mainly since the late Oligocene. Results from the present study provide insights into the historical assembly of herbaceous flora in northern temperate regions.
... Arenaria Linnaeus (1753: 423) (Caryophyllaceae Juss., Alsinoideae Fenzl, Arenarieae Kitt.) is a genus comprising about 160-175 species of annual and perennial herbs mostly distributed in the northern temperate and subartic regions, the Mediterranean, Mexico, and the Andes of South America (see Bittrich 1993, Hernández-Ledesma et al. 2015, POWO 2022. The molecular data by Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014), and Sadeghian et al. (2015) showed that Arenaria is polyphyletic and several names were trasferred to other genera (see e.g., Conti et al. 2014, Dillenberger & Kadereit 2014, Sadeghian et al. 2015, Iamonico 2016. In addition to taxonomic issues, there are several nomenclatural questions remaining to be addressed for names that were previously published elsewhere (see e.g., POWO 2022a), and several of them were not typified (see e.g., Iamonico 2013Iamonico , 2014Iamonico , 2016Iamonico , 2019Iamonico , 2022. ...
... Arenaria Linnaeus (1753: 423) (Caryophyllaceae Juss., Alsinoideae Fenzl, Arenarieae Kitt.) is a genus comprising about 160-175 species of annual and perennial herbs mostly distributed in the northern temperate and subartic regions, the Mediterranean, Mexico, and the Andes of South America (see Bittrich 1993, Hernández-Ledesma et al. 2015, POWO 2022. The molecular data by Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014), and Sadeghian et al. (2015) showed that Arenaria is polyphyletic and several names were trasferred to other genera (see e.g., Conti et al. 2014, Dillenberger & Kadereit 2014, Sadeghian et al. 2015, Iamonico 2016. In addition to taxonomic issues, there are several nomenclatural questions remaining to be addressed for names that were previously published elsewhere (see e.g., POWO 2022a), and several of them were not typified (see e.g., Iamonico 2013Iamonico , 2014Iamonico , 2016Iamonico , 2019Iamonico , 2022. ...
Article
The nomenclature of the names Alsine weberbaueri, Arenaria musciformis, and Scleranthus peruvianus is investigated. The names were typified on specimens preserved at HOXA, MOL, and P. All these names are to be included in Arenaria. Concerning Alsine weberbaueri, a new combination (Arenaria weberbaueri) is proposed, whereas for Arenaria humboldtiana, a new name is provided for Scleranthus peruvianus since a previous Arenaria peruviana by Poeppig ex Rohrbach was validly published (Art. 53.1 of ICN).
... The genus Arenaria L. (Alsinoideae Fenzl, Caryophyllaceae Juss.) comprises about 160 species of annual and perennial herbs mostly distributed in the northern temperate regions, the Mediterranean, and the Andes of South America (Hernández-Ledesma et al., 2015 and literature therein). The molecular data by Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014), and Sadeghian et al. (2015) confirmed that Arenaria is polyphyletic. As a result, several species were included into other genera, such as Ermogone Fenzl, Honckenya Ehrh., Minuartia L., Moheringia L., Odontostemma Benth ex G. Don, Sabulina Rchb., Solitaria (McNeill) Sadeghian & Zarre, and Spergularia (Pers.) ...
... & C. Presl. (Conti et al., 2014;Dillenberger & Kadereit, 2014;Iamonico, 2014). The Flora of Chile comprises eight Arenaria species, of which three are endemic (Ulloa Ulloa et al., 2017;Rodriguez et al., 2018: 253-254;Zuloaga et al., 2019 [app. ...
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A nomenclatural study of the Arenaria names referring to the species found in Chile is presented. Three names, A. fastigiata, A. multicaulis, and A. scopulorum, are illegitimate (Arts. 53.1 and 52.1 of ICN); a new name is proposed for A. fastigiata and A. multicaulis; A. scopulorum was also nomenclaturally superfluous including the type of A. digyna (Art. 52.1 of the ICN). Arenaria digyna, A. microphylla, A. oligosperma, A. pleurantha, and A. rivularis are lectotypified, based on specimens preserved at E, HAL, K, P, and SGO (isolectotypes at B, K, P, and SGO). One epitype (at SI) was designated for A. rivularis.
... 1A, 1B). Notably, in a redefined circumscription of Minuartia, Dillenberger and Kadereit (2014) not only accepted Eremogone as a distinct genus, but to make it monophyletic they suggested its expansion by proposed transfer of the members of Minuartia subgen. Spergella (Fenzl) McNeill to Eremogone. ...
... The fact that the said phylogenetic analysis included two Himalayan members of subgenus Eremogoneastrum, viz., Arenaria bryophylla and A. kansuensis, both falling in the distinct Eremogone clade, strongly support the placement of Himalayan members of Eremogoneastrum in the genus Eremogone instead of Arenaria, s.s. Following these results, Dillenberger and Kadereit (2014) (Hara andTebbs 1979, Singh andDiwakar 2010). Of these, only one, A. polytrichoides, is reported from Indian western Himalaya. ...
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The genus Arenaria L. (Caryophyllaceae) in the Indian Western Himalaya is studied in detail and rearranged. Three genera, namely Dolophragma Fenzl, Eremogone Fenzl and Odontostemma Benth. ex G. Don, which were previously treated as subgenera under the genus Arenaria, are here recognized as distinct genera and corresponding species of Arenaria are transferred to them. As concluded in phylogenetic studies, subgenus Eremogoneastrum Fenzl is treated as a part of the genus Eremogone and new combinations are proposed for eight western Himalayan taxa transferred here under the genus Eremogone. Species hitherto treated in Arenaria subgenus Arenaria are retained as it is, except the sole representative of section Compressae McNeill, which is shifted under a newly described monotypic genus Himgiria. Similarly, three species hitherto under Arenaria subgenus Solitaria McNeill now form part of newly described Sino-Himalayan genus Shivparvatia. A key to the genera of Arenaria and its allies reported from Indian western Himalaya is also provided.
... The family is monophyletic, as circumscribed by Bittrich [4], but the traditional classification into three subfamilies [4,5], which is based on stipule, petal, sepal, and fruit features, does not provide monophyletic groups and should be replaced with the tribe-based scheme [1,2]. According to The Caryophyllales Network [6], 11 According to phylogenetic analyses of the whole family [1, 2,7], the genus Drymaria Willd. ex Schultes belongs to the tribe Polycarpeae, a well-supported clade which is basal to a large group including many other tribes of Caryophyllaceae, i.e., Sperguleae, Sclerantheae/Sagineae, and Ermogoneae/Caryophylleae/Sileneae. ...
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A new species of Drymaria from the Kohima District (Nagaland of North-East India) is described and illustrated based on both molecular data (the ITS region of nuclear DNA) and morphometric analyses (clustering, PCA, DA, and MANOVA). The new species resembles D. cordata but differs with respect to the shape of the sepal (oblong, incurved at the margin vs. lanceolate, not incurved at the margin), petals (oblong or linear vs. ovate–lanceolate), and bract (foliaceous, glabrous, non-prominent vs. non-foliaceous, pubescent, and prominent). For nomenclatural purposes, the typification of Drymaria villosa by Duke (in 1961) was corrected according to Art. 9.10 of the Shenzhen Code.
... Traditional Stellaria (i.e., Stellaria s.l.) shows high morphological diversity, and all the previous studies based on both morphological and molecular evidence revealed that Stellaria s.l. has a complex evolutionary history (Smissen et al. 2002, Chen et al. 2010, Greenberg & Donoghue 2011, Sharples & Tripp 2019. Harbaugh et al. (2010) revealed Stellaria s.l. as paraphyletic, and the finding was supported by Fior et al. (2006), Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014) and Zhang et al. (2017). Recent studies supported the segregation or resurrection of several genera, i.e., Adenonema Bunge (1835: 548), Engellaria Iamonico (2021: 4), Hesperostellaria Gang Yao, B. Xue & Z.Q.Song (2023: 15), Maguirellaria Iamonico (2023: 240), Mesostemma Vvedensky (1941: 4), Nubelaria M.T. Sharples & E.A.Tripp (2019: 869), Rabelera M.T. Sharples & E.A.Tripp (2019: 870), Reniostellaria Gang Yao, B. Xue & Z.Q.Song (2023: 15), Schizotechium (Fenzl) Reichenbach ex Kafle & G.Parmar (2024: 30) and Torreyostellaria Gang Yao, B. Xue & Z.Q.Song (2023: 15) from Stellaria s.l. to delimit the core Stellaria (i.e., Stellaria s.s.) in the tribe Alsineae (Pusalkar & Srivastava 2016, Sharples & Tripp 2019, Arabi et al. 2022, Xue et al. 2023. ...
Article
Based on distinctive morphology, Stellaria rajbhandarii is described as a new species from Nepal, and S. neotomentosa is resurrected since it is a species distinct from S. nepalensis. This study has provided the diagnosis as well as a complete description, phenology, distribution, habitat, and photoplates of both species.
... Harbaugh et al. (2010), suggesting that traditional subfamilies do not reflect natural groups, studied matK, trnL-F and rps16 chloropslast gene sequences and proposed a new classification system, replacing the traditional classification of 3 subfamilies, and divided the family Caryophyllaceae into 11 tribes including the new tribe Eremogoneae Rabeler W.L.Wagner. The phylogenetic position of Eremogone, which is placed outside of Arenaria and within a separate tribe called Eremogoneae, has been extensively studied (Greenberg & Donoghue 2011, Harbaugh et al. 2010, Dillenberger & Kadereit 2014, Sadeghian et al. 2015. As a result of phylogenetic studies, the genus Eremogone was formed by collecting species in Arenaria subgenus Eremogone Fenzl (approximately 70 species), Arenaria subgenus Eremogoneastrum F.N.Williams (approximately 22 species) and Minuartia subgenus Spergella (Fenzl) McNeill (approximately 3 species). ...
Article
The palynological characters of 17 taxa belonging to genus Eremogone Fenzl (Caryophyllaceae) from Türkiye were examined using light microscopy and scanning electron microscopy. Eremogone pollen in Türkiye is found to have an average size of 31.31 μm (26.62‒36.65 μm), pore sizes ranging from 6.52 to 9.24 μm, inter-pore distances ranging from 9.07 to 12.57 μm, pore numbers varying from 5 to 14 and 5‒30 nanoechini distributed in an area of 10 × 10 µm. The ornamentation was identified as nanoperforate, nanoechinate, or nanofaveolate surface elements in various combinations. Clustering analysis and Principal Component Analyses do not show clear groups, although more similar taxa were brought together through the analysis of pollen micromorphological data rather, the nanoechinate base width, nanoechinate length, and pore diameter length are the most valuable variables for separating the examined taxa.
... The pattern of a polyploid Arctic-alpine distribution with an overlap into the Balkan Peninsula and the Mediterranean area is perfectly The taxonomy of the group is still under dispute, and the number of taxa and their delimitation strongly differs between various concepts (Graebner et al., 1918;Hayek, 1922;Pawłowski, 1939;Halliday, 1964Halliday, , 1993Kamari and Constantinidis, 1994;Dvořáková, 2000). Although Dillenberger and Kadereit (2014) included several accessions of this group into the phylogenetic reconstruction of Minuartia s.l. and Nunvářová Kabátová et al. (2019) examined plastid DNA variation, the evolutionary history and relationships among its taxa still remain largely unresolved. ...
Article
One of the major goals of contemporary evolutionary biology is to elucidate the relative roles of allopatric and ecological differentiation and polyploidy in speciation. In this study, we address the taxonomically intricate Sabulina verna group, which has a disjunct Arctic–alpine postglacial range in Europe and occupies a broad range of ecological niches, including substrates toxic to plants. Using genome-wide ddRAD sequencing combined with morphometric analyses based on extensive sampling of 111 natural populations, we aimed to disentangle internal evolutionary relationships and examine their correspondence with the pronounced edaphic and ploidy diversity within the group. We identified two spatially distinct groups of diploids: a widespread Arctic–alpine group and a spatially restricted yet diverse Balkan group. Most tetraploids exhibited a considerably admixed ancestry derived from both these groups, suggesting their allopolyploid origin. Four genetic clusters in congruence with geography and mostly supported by morphological traits were recognized in the diploid Arctic–alpine group. Tetraploids are split into two distinct and geographically vicariant groups, indicating their repeated polytopic origin. Furthermore, our results also revealed at least five-fold parallel colonization of toxic substrates (serpentine and metalliferous), altogether demonstrating a complex interaction between geography, challenging substrates and polyploidy in the evolution of the group. Finally, we propose a new taxonomic treatment of this complex.
... Having a clear understanding of the nomenclature and the changes it has undergone in the past is also key to the legal protection of any rare species (Mace, 2004), including plants (Standley, 1992;Schatz, 2002). For example, if an SSA was to be developed for Minuartia patula, which has undergone multiple changes of nomenclature and delimitation (Kartesz, 1994;Kartesz, 1999;Dillenberger and Kadereit, 2014;Schilling et al., 2022), under which sense [or taxon concept (Franz et al., 2008;Sigovini et al., 2016)] would it be recognized? In cases such as these, consulting herbarium specimens, taxonomic databases and literature, or even taxonomic experts to determine the taxonomic unit of conservation is prudent. ...
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Funding for rare plant conservation is limited. In addition, many aspects of the biology and ecology of rare plants are unknown. Therefore, low-cost data generation approaches to fill these gaps should be pursued. Herbarium specimens can be used as a low-cost alternative to learn about the basic biology and ecology of rare plant species. The information provided on herbarium labels has dramatically increased in recent decades to include precise locality (i.e., latitude/longitude), exact dates, habitat, associated species, and substrate. In addition, herbarium specimens are being digitized and the resulting images and data are available via clearinghouses such as GBIF and SEINet. Already, herbarium specimens of rare plants have been used to develop habitat suitability models, predict range shifts, and assess changes in flower phenology due to climate change. Herbarium specimens can also provide a wealth of information about the reproductive biology and biotic interactions of rare plants. In this paper, we will demonstrate how this information can be accessed and present a practical application for using this information to populate an important federal listing document in the USA, Species Status Assessments (SSA). We will provide examples from the literature, as well as case studies from our own research, to demonstrate how this information can be collected from herbarium specimens and how and where to incorporate this information into SSAs. More generally, data gleaned from herbarium specimens can become part of a conservationist’s tool kit to further our knowledge of past, present, and future trends for rare plants. Additional knowledge of a species’ biology and ecology allows land managers and conservationists to make more informed decisions and allows for greater protection of listed species.
... Schischkin and M. mediterranea (Link) K. Malý: some recent studies, corroborated by molecular data (e.g. Dillenberger & Kadereit 2014), place these two species in a separate genus, Sabulina Reichenb., a point of view already followed by some contemporary western European Floras, e.g. H24 and Stace (2019). ...
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Following the publication of the seventh edition of the Nouvelle Flore, in 2023, this contribution provides an overview of nomenclatural and taxonomic changes compared to the previous edition of the Flora, which was published in 2012.
... Stellaria L. (Caryophyllaceae Juss.) is a genus traditionally including 150-200 species occurring in the temperate regions of Eurasia and North America and at the higher altitudes of tropical areas, being mostly diverse in the mountains of central Asia (Hernández-Ledesma et al. 2015, Tikhomirov 2016. Recent molecular studies (Greenberg & Donoghue 2011, Dillenberger & Kadereit 2014, Sharples & Tripp 2019 have shown that this genus is polyphyletic and several taxa have been shown to belong to other genera, e.g., Adenonema Bunge, Cerastium L., Hartmaniella M.L.Zhang & Rabeler, Mesostemma Vved., Minuartia L., Mononeuria Rchb., Nubelaria Sharples & Tripp, Pseudocherleria Dillenberg. & Kadereit, and Rabelera Sharples & Tripp. ...
Article
Molecular analyses have shown that Stellaria is a polyphyletic genus and several species are closely related to other genera of Caryophyllaceae. Among the excluded taxa, a group of four species (S. antillana, S. howardii, S. minutifolia, and S. ovata) appear to be not included in the core of Stellaria and are possibly related to the genus Triplateia. Despite this molecular evidence, as well as the morphology of these four species, a formal proposal for a new taxon is still lacking. I here describe the genus Maguirellaria gen. nov., including Maguire’s Stellaria howardii and S. minutifolia. S. antillana and S. ovata, which display a very different morphology in comparison with Maguire’s species, were never considered in the same molecular study; therefore, I prefer to avoid a taxonomic decision for the moment.
... The family, the biodiversity center is the Eastern Mediterranean and Irano-Turanian region, is mainly distributed in northern temperate, mountainous and alpine regions (POWO 2023). The monophyly of the family was supported by molecular evidences (see e.g., Greenberg & Donogue, 2011), but its classification is still partially unresolved, and several genera need further studies being critical from both taxonomical and nomenclatural points of views (see e.g., Dillenberger & Kadereit 2014, Iamonico 2021a, 2021b, 2022, Iamonico & Domina 2015, De Luca et al. 2022, El Mokni et al. 2023, Montesinos-Tubeé & Iamonico 2023 and need further studies, among these genera, Gypsophila Linnaeus (1753: 406). ...
Article
Gypsophila dumanii, a new species belonging to the section Capituliformes, is here described and compared with the most similar G. olympica. A detailed description, original photographs, the current distribution area, and the conservation status of G. dumanii are given.
... The genus Stellaria L. (Caryophyllaceae) comprises 150-200 species mainly occurring in the temperate regions of Eurasia, North America, and at the higher altitudes of tropical areas (see e.g., Chen & Rabeler 2001, Hernández-Ledesma et al. 2015, Xu & Ma 2018, Wang et al. 2020a, 2020b, Xu et al. 2020. Molecular studies (Greenberg & Donoghue 2011, Dillenberger & Kadereit 2014, Sharples & Tripp 2019 Iamonico], based on the above-mentioned molecular data as well as on taxonomic considerations. After these changes, according to Sharples & Tripp (2019) and Iamonico (2021), Stellaria would now consist of 111 species, but a worldwide revision of Stellaria is still lacking and most of the taxonomic data can be found in comprehensive floras [see e.g., Chater & Heywood (1993) for Europe, Chen & Rabeler (2001) for China, Morton (2005) for North America, Miller & West (2012) for Australia]. ...
Article
Stellaria ruderalis and S. cupaniana (Caryophyllaceae, Alsinoideae, Alsineae) are recorded from Tunisia for the first time, S. ruderalis being also new to continental Africa. Descriptions, notes on habitat, ecology, and distribution in Tunisia are given. Original photographs together with a key to the Stellaria taxa of Tunisia are also provided. The typification of the name Alsine cupaniana is discussed and a specimen preserved at LY is designated as the lectotype.
... Οι τρεις παραδοσιακές υποοικογένειες Silenoideae, Alsinoideae και Paronychioideae (Pax, 1899;Bittrich, 1993) δεν φαίνεται να επιβεβαιώνονται από τα ευρήματα της μοριακής φυλογενετικής, επομένως έχουν εγκαταληφθεί στις πιο πρόσφατες θεωρήσεις της οικογένειας, αντικαθιστώμενες από 11 φυλές (tribes) συγγενικών γενών (Ηarbaugh et al., 2010;Greenberg & Donoghue, 2011). Αντιστοίχως, σε επίπεδο γενών αναμένονται αρκετές ανακατάταξεις τα επόμενα χρόνια χάρη στην ευρύτερη και πιο ευχερή χρήση των μοριακών δεικτών, με χαρακτηριστικό παράδειγμα τη διάσπαση του πολυφυλετικού γένους Minuartia σε 11 γένη (Dillenberger & Kadereit, 2014). ...
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Dianthus cruentus Griseb. (Caryophyllaceae) is a herbaceous perennial species native to the Balkan Peninsula, including Greece, and western Asia. It is a species of great ornamental interest due to its architectural inflorescences and its compact blue-green foliage, particularly resistant to dry and poor soils and attractive to pollinators such as butterflies and bees. In addition, it is a species of medical potential due to its strong antioxidant action. In the present study, the determination of the appropriate conditions for maximizing the germination of its seeds in both in vitro and in vivo conditions was initially investigated. Two seed lots collected from a native population of the species on Mount Kallidromo, Fthiotida, dry-stored for 0 or 12 months, were used, without any pre-treatment. Regarding in vitro germination, the seeds were surface sterilized with a 20% commercial bleach water solution for 10 minutes, were then rinsed with distilled water and afterwards were placed in Petri dishes containing ½ MS medium, under 16 h photoperiod from 37.5 μmol m-2 s-1 fluorescent light, at temperatures of 5, 10, 15, 20 or 25 °C. The optimum temperature for seed germination was 15 °C, in which the highest germination ability in both recently collected (98%) and 12-month-old seeds (100 %) was observed. The lowest germination percentages were recorded at the two extreme temperatures of 5 °C and 25 °C, with particularly low values for recently collected seeds (18% and 15%). At all temperatures, minus that of 5 °C, the seeds germinated rapidly (T50 = 2-6 d). The duration of the seeds’ dry storage significantly affected their germination, increasing the germination percentage at the 2 extreme temperatures and demonstrating the positive effect of dry afterripening. As for ex vitro germination, the seeds were placed in pots with a mixture of peat-perlite (1: 1, v/v) and were incubated in growing chambers at temperatures of 15, 20 or 25 °C. The highest germination percentage was observed at 15 °C for both seed lots (97-99%), while the lowest at the temperature of 25 °C (62% and 4% for the 0- and 12-months-old seeds respectively). Germination ability was slightly lower in ex vitro germination, especially at 25 °C. Furthermore, the micropropagation of the species from explants excised from 2-month-old seedlings was investigated on solid substrates. Substrates containing MS medium and 8 g L-1 agar were used in almost all substages of the proliferation stage, and the effects of cytokinins BA, 2iP and ZEA, with the absence or the presence of auxin NAA, were investigated, using explants derived from non-hyperhydric or hyperhydric shoots. In the initial culture substrates containing 0.1 mg L-1 BA or 0.1 or 0.5 mg L-1 2iP the explants produced shoots at a high percentage (72-87%), but many of these were shown to be hyperhydric, with abnormal morphology and vitreous appearance. The number of hyperhydric shoots and the number of nodes seemed to be higher on substrates containing 2iP, with the largest number of normal shoots (2.3) appearing in the substrate containing 0.1 mg L-1 BA. During the 1st subculture stage, hyperhydric shoots were observed in bigger numbers in the substrate containing 2iP, while an increase in shoot proliferation and the number of hyperhydric shoots was correlated with the use of hyperhydric explants grown on hormone-free MS substrate. In the 2nd subculture stage, the presence of 0.1 mg L-1 BA yielded the largest number of shoots regardless of the physiology of the explants, while the addition of 0.05 mg L-1 resulted in a significant increase in the number of normal shoots and a reduction in hyperhydricity rates when hyperhydric explants were used. In the 3rd subculture, when explants derived from nonhyperhydric shoots were used the highest value of the multiplication index (4.5) and the maximum number of shoots (3.6) was recorded in MS substrate containing 2 mg L-1 BA. Increasing the concentration of agar to 12 g L-1 produced the largest number of shoots (2.4) when hyperhydric explants were used. In both cases, the highest percentages of hyperhydricity were recorded on MS substrate containing 0.1 mg L-1 ZEA. In contrast, during the 4th subculture, the use of this substrate resulted in the greatest multiplication index value at this stage (4.8). Overall, the use of MS substrate with 0.1 mg L-1 BA and 0.05 mg L-1 NAA resulted in the highest values of the proliferation index (5.1) regardless of the physiology of the explants used, with its maximum value (5.2) recorded from the culture of hyperhydric explants on MS substrate containing 0.1 mg L-1 BA and 12 g L-1 agar. The presence of phytohormones in the substrates increased the production of callus around the base of the explants, while high concentrations of cytokinins lessened the occurence of roots. D. cruentus microshoots were rooted in MS or ½ MS substrates in the presence or absence of IBA. The use of MS substrate resulted in lower rooting rates (55-70%) when lower concentrations of IBA were used (0 or 0.1 mg L-1). The optimal concentration of IBA was found to be 0.5 mg L-1 regardless of the multiplication substrate where the microshoots originated or the nutrient medium used in the rooting substrate, with rooting rates of 83-100%, while in some instances a carry over effect, caused by the cytokinin content of the shoot proliferation substrates, was observed on the roots’ number and median length. The ex vitro acclimatization of the rooted microshoots was completed with a success rate of 91% using a peat:perlite substrate (1:1, v/v). Significant morphological differences were observed between acclimatized plantlets, including differences in stem length and total number of shoots and nodes, which were correlated with the influence of different shoot proliferation and/or rooting substrates, while the presence of IBA during rooting seemed to have an amendable effect on their survival. Finally, the use of an integrated, 60-days-long propagation protocol with shoot proliferation and rooting occurring concurrently in one stage on MS substrates free of PGRs or containing BA or 2iP and NAA produced rooted young plants at a high percentage (88%), which had the maximum observed total number of shoots and nodes and were then successfully acclimatized, surviving at a percentage of 86%. Scientific area: Horticulture Keywords: Dianthus cruentus, native ornamental plant, germination ecophysiology, seed germination in vitro, seed germination temperature, seed lot age, dry afterripening, in vitro propagation, solid culture, seedling explant, hyperhydricity, hyperhydric explant, callus induction, in vitro rooting, ex vitro acclimatization, cytokinins, IBA, carry over effect, acclimatized plantlets’ morphology
... aus Minuartia s. l. ausgegliedert werden. Dillenberger & Kadereit (2014) konnten zeigen, dass die verwendeten Merkmale zur Gattungsabgrenzung von Minuartia völlig ungeeignet waren. Die mehrfache Ausschlüsselung von Minuartia im Hauptschlüssel der Familie hat schon in früheren Auflagen des Grundbandes gezeigt, dass Minuartia s. l. morphologisch divers ist und sich schwer zusammenfassen lässt, jedoch die Aufteilung in vier natürliche Gattungen erleichtert. ...
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The 22nd edition of the Rothmaler field flora was published in 2021, and the present article gives comments on the numerous, especial major changes. We added new species, and for the first time included an Algae group; the stoneworts, Characeae, which in many practical respects resemble vascular plants. For all species, biological data were revised, and chromosome numbers were added if available for accessions from Germany. Notes on distribution within Germany were completely restructured, employing new physico-geographical rather than political mapping units, and using data from the current distribution atlas of vascular plants in Germany. Taxonomy and nomenclature in the Rothmaler largely follow the standard checklist of vascular plants in Germany; deviations are explained and justified. Die 22. Auflage des Gundbandes der Rothmaler Exkursionsflora ist 2021 erschienen und der vorliegende Artikel gibt Kommentare und Hinweise zu vielen, vorwiegend größeren Neuerungen. Wir haben im Grundband nicht nur neue Taxa aufgenommen, sondern mit den Armleuchteralgen, den Characeae, auch erstmals eine Algengruppe verschlüsselt, die aber in vieler, insbesondere ökologisch-praktischer Hinsicht durchaus den Gefäßpflanzen ähnelt. Für alle Arten im Band wurden die biologischen Angaben durchgesehen; sofern Daten zu Akzessionen aus Deutschland verfügbar waren, wurde Chromosomenzahlen neu eingefügt. Die Angaben zur Verbreitung in Deutschland wurden basierend auf dem Deutschlandatlas neu zusammengestellt, wobei jetzt stärker naturräumliche als politische Verbreitungseinheiten genutzt wurden. Taxonomie und Nomenklatur im Grundband folgen weitgehend der Florenliste von Deutschland, Abweichungen werden hier diskutiert und ggf. begründet.
... & J.Presl including about 3000 species (100 genera) which are primarily distributed in the Holarctic region, having a center of diversity in the Mediterranean and Irano-Turanian regions (Bittrich 1993, Heywood 1998, Hernández-Ledesma et al. 2015. Greenberg & Donoghue (2011) demonstrated the monophyly of Caryophyllaceae using molecular data, but its classification remains partially unresolved since several genera are critical from the taxonomical and nomenclatural point of view (Conti et al. 2014, Dillenberger & Kadereit 2014, Iamonico 2015, 2021a, 2021b, Iamonico & Domina 2015. ...
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The seed and leaf structures of some species belonging to the genus Saponaria (Caryophyllaceae) were examined using SEM. These species were: S. calabrica, S. dalmasii (endemic), S. glutinosa, S. halophila (endemic), S. karapinarensis (endemic), S. kotschyi (endemic), S. mesogitana, S. officinalis, S. orientalis, S. pamphylica (endemic), S. picta (endemic), S. pinetorum var. elatior (endemic), S. prostrata subsp. anatolica (endemic), S. prostrata subsp. calvertii, S. prostrata subsp. prostrata (endemic), S. pumilio, and S. tridentata. The results obtained show that the micromorphology of the seed are useful to distinguish the studied species. Four groups can be distinguished using the number of seed per capsule. In addition, the shape of the seed, microsculpture, shape and margin of testa cells were found as important characteristics to identify the various species. Leaf anatomy reveals that trichome type and mesophyll symmetry define significant differences between the studied species.
... Augustea Iamonico, Facchinia Rchb., Mcneillia Dillenb. & Kadereit, Sagina L., Scleranthus L.;Smissen et al., 2003;Dillenberger & Kadereit, 2014;Iamonico, 2015Iamonico, , 2016 y, en algunos casos, merecen aclaraciones sobre su nomenclatura y taxonomía. Tal es el caso de Cardionema DC., un género nativo del Nuevo Mundo, con unas 6 especies, distribuido desde los Estados Unidos de América hasta el sur de la Patagonia (Hartman, 2005;Ulloa Ulloa et al., 2017;Hernández-Ledesma et al., 2015). ...
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The genus Cardionema (Caryophyllaceae) is native to the New World and it comprises six species of perennial herbs, with spiny leaves and sepals, membranous stipules and bracts, and reduced petals. This study aims to update the richness, morphology and distribution of Cardionema in Argentina, based on the study of herbarium specimens. Four species of Cardionema are accepted for the Flora of Argentina; a description of each taxon, an identification key, illustrations and pictures, a comparative-morphology based table, and distribution maps are included. Two lectotypes are also designated.
... Molecular studies on Petrorhagia, Velezia and Gypsophila revealed the relationship among related taxa (Korkmaz & Doğan 2015;Poyraz et al. 2012;Hilooğlu et al. 2016). Molecular phylogenetic studies based on ITS and plastid trnQ-rps16 DNA in the largest genera (Gypsophila L., Dianthus L., Arenaria L., Minuartia L. and Silene L.) have resulted in revised treatments including new genera and synonymy (Dillenberger & Kadereit 2014;Madhani et al. 2018;Sadeghian et al. 2015). ...
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Gypsophila malyerii Hamzaoğlu & Koç, a new species of sect. Capituliformes, is described and illustrated from Turkey. Information on distribution, habitat and conservation status are given. The most similar species is G. osmangaziensis. The morphology and micromorphology of seed and pollen characters of the two closely related species are compared.
... Six out of these nine latter species were later recognized as part of the following two other genera: Mononeuria Rchb. Dillenberger and Kadereit, 2014) which is sister to some Minuartia members. They need further study and are part of an ongoing study by the author of the present paper (Iamonico, in prep.). ...
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Background and Aims: Stellaria traditionally comprises 150-200 species, mainly distributed in the temperate regions of Eurasia and North America. Molecular studies demonstrated that Stellaria is polyphyletic and includes about 120 species. The genus has a high phenotypic variability which has led to nomenclatural disorders, making the identification of the various species difficult. A note is presented about a taxon currently accepted under the genus Stellaria -Stellaria obtusa- which should be recognized as a separate genus, here proposed as Engellaria gen. nov. Methods: This study is based on examination of specimens of American and European herbaria and analysis of relevant literature. Key results: Available molecular data show that Stellaria obtusa is not included in the Stellaria s.s. clade, but instead is basal to another clade comprising the genera Honckenya, Schiedea, and Wilhelmsia. Stellaria obtusa was, therefore, compared with these three groups and with morphologically similar apetalous members of Stellaria s.s. (S. crispa, S. media, S. pallida, and S. irrigua). The results obtained lead to the recognition of S. obtusa as a separate new North American monotypic genus. A diagnostic key of the apetalous members belonging to the American Caryophyllaceae genera is proposed. Finally, the names Stellaria obtusa and S. washingtoniana (= S. obtusa) are lectotypified based on specimens deposited, respectively, at UC (isolectotypes at GH, NY, and YU) and GH (isolectotypes at BM, CAN, CAS, CS, DOV, F, GH, K, MIN, MSC, NY, US, and VT). For the name Alsine viridula (= S. obtusa) the holotype was found at US (isotypes at CAS, F, GH, NY, OSC, RM, and UC). Conclusions: Stellaria obtusa does not belong to the genus Stellaria. The present study shows that the combined use of morphological data and phylogenetic analyses helped to clarify the taxonomic position of difficult plant groups, as in Stellaria.
... Given the dynamic range history of plant taxa in the EAS, we here examine the role of hybridization in the evolution of a subclade of Cherleria L. (formerly Minuartia L., Caryophyllaceae; [21,54]) endemic to the EAS (Fig. 1). Cherleria as a whole contains 19 species and has a circumboreal distribution, with incursions into the high mountains of Eurasia and North America. ...
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Background Cherleria (Caryophyllaceae) is a circumboreal genus that also occurs in the high mountains of the northern hemisphere. In this study, we focus on a clade that diversified in the European High Mountains, which was identified using nuclear ribosomal (nrDNA) sequence data in a previous study. With the nrDNA data, all but one species was monophyletic, with little sequence variation within most species. Here, we use genotyping by sequencing (GBS) data to determine whether the nrDNA data showed the full picture of the evolution in the genomes of these species. Results The overall relationships found with the GBS data were congruent with those from the nrDNA study. Most of the species were still monophyletic and many of the same subclades were recovered, including a clade of three narrow endemic species from Greece and a clade of largely calcifuge species. The GBS data provided additional resolution within the two species with the best sampling, C. langii and C. laricifolia , with structure that was congruent with geography. In addition, the GBS data showed significant hybridization between several species, including species whose ranges did not currently overlap. Conclusions The hybridization led us to hypothesize that lineages came in contact on the Balkan Peninsula after they diverged, even when those lineages are no longer present on the Balkan Peninsula. Hybridization may also have helped lineages expand their niches to colonize new substrates and different areas. Not only do genome-wide data provide increased phylogenetic resolution of difficult nodes, they also give evidence for a more complex evolutionary history than what can be depicted by a simple, branching phylogeny.
... or Eken et al. (2006), where area-based lists of selected rare species are offered. The naming of the species follows largelyGüner et al. (2012), with the exception of the following taxa (references to the taxonomic backbones in brackets): Anemonastrum, Asteraceae, Lomelosia, Noccaea, Odontarrhena and pteridophytes (The Euro+Med PlantBase [http://ww2.bgbm.org/EuroPlusMed/query. asp]; Caryophyllales(Dillenberger and Kadereit 2014;Hernández-Ledesma et al. 2015;Madhani et al. 2018), Lamium ...
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Abstract. This chapter characterizes the flora and vegetation of the North-Western Caucasus, taking the Karachaevo-Cherkessian Republic (a mountainous region in the southwestern Russia) and the Teberda State Biosphere Reserve (a protected nature area with restricted access located on the northern spurs of the Greater Caucasus Range) as examples. The Teberda State Biosphere Reserve is located in the southern part of the Karachaevo-Cherkessian Republic at the upper reaches of the Teberda and Kizgych rivers and their tributaries. The elevation in the Teberda State Biosphere Reserve varies from 1259 to 4046 m a.s.l. The treeline runs at 1700–2400 m a.s.l., and the upper limit of vascular plants is found at 3750 m a.s.l. About 83% of the Reserve area lies above 2000 m a.s.l. Rock and scree outcrops occupy 26% of the Reserve. The wide elevational range, complex topography, and sharp changes of microclimatic conditions determine a high species diversity and variety of plant communities. The flora of the Karachaevo-Cherkessian Republic currently comprises 1959 species of vascular plants. There are 10 floristic elements in the indigenous flora of the Republic, i.e. Holarctic, Palaearctic, Panboreal, Euro-Mediterranean, Mediterranean, Ancient Mediterranean, Irano-Turanian, Pontic-South- Siberian, Pluriregional, and Endemic. A total of 343 species (17.5%) are endemics. Six elevational belts (steppe, deciduous broad-leaved forest, coniferous forest, subalpine, alpine, and subnival) and major vegetation types, including steppes, deciduous and coniferous forests, crooked-stem elfin woods, tall grass communities, fens, alpine and subalpine meadows, snowbeds, lichen heaths, pioneer communities on rocks, screes and talus slopes are reviewed. The Caucasus is considered a biodiversity hotspot with significant levels of endemism at both species and ecosystem levels. Currently it is prioritized as one of the most vulnerable regions with a high irreplaceability.
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In this paper, we reviewed the records of Caryophyllaceae family of flowering plants reported from Uttarakhand state of India. Family Caryophyllaceae Juss. in Uttarakhand is represented by 24 genera 75 species, 2 subspecies and 6 varieties. Based on the herbarium history, literature survey and additional observations by the authors, 11 taxa were excluded from the flora of Caryophyllaceae of Uttarakhand. Cherleria sedoides was reported for the first time from India and Uttarakhand. Hitherto, the misidentification of the Cherleria sedoides and Cherleria biflora in the Himalayas, and identification ambiguity between Cerastium glomeratum and C. fontanum subsp. vulgare were resolved. We found that Stellaria depressa is a new record to the flora of Uttarakhand. New combinations of Stellaria patens subsp. patens, Stellaria patens subsp. semivestita and Sabulina kashmirica var. foliosa have been proposed. Based on the taxonomic history, and observations of the live and herbarium specimens, the synonymy of Sagina purii for S. apetala was concluded. This study summarises and presents the updated flora of Caryophyllaceae family in Uttarakhand based on these new findings.
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Molecular phylogeny has gained a crucial role for plant taxonomy in recent years. In some cases sequence comparison of the chloroplasts or nuclear genes and even whole genomes led to novel and unexpected results in taxonomic positions. For example, after publication of phylogenetic data a genus Minuartia s. l. was considered to be a paraphyletic group and subjected to taxonomic revision according to cladistics concepts. As a result, several new genera were identified and some genera were restored including Cherleria L., which includes 19 species. The genus Cherleria is of particular interest due to significant habitual differences in its representatives. Thus, to clarify the relationships between intergeneric taxa a search for new taxonomically significant traits is needed. For these purposes, the biomorphological characteristics of representatives of six model species of Cherleria were studied. In the article the most significant biomorphological characteristics such as the model of shoot formation, the morphology of the cushion, the nature of shoot branching, the presence or absence of specialized vegetative shoots for capturing space, the growth strategy and the structure of the monocarpic shoot are proposed to aid for identification of taxonomic position. Analysis of the biomorphological features of the studied species identified four growth strategies: a) the strategy of centrifugal growth; b) the strategy of centrifugal growth with the rare appearance of radially directed lateral specialized shoots of the semi-rosette type; c) the strategy of vector growth by active branching at the early stages of development, with subsequent “fading” in formation of lateral axes; d) the strategy of vector growth with the help of semi-rosette plagiotropic shoots with the subsequent filling of the internal space of the cushion due to orthotropic rosette shoots. Comparison of molecular phylogeny and biomorphology data allows us to identify biomorphological apomorphies: monopodial plagiotropic-semi-rosette–orthotropic-rosette model of shoot formation with specialized semi-rosette shoots for capturing territory, the presence of dicyclic monocarpic shoots, as well as plesiomorphies: monopodial rosette model of shoot formation, formation of a densely filled cushion, monocyclic, simply arranged monocarpic shoots. Based on these data, we put forward an assumption that for the genus Cherleria the initial model of shoot formation was the monopodial rosette model.
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The phylogenetic studies of the tribe Alsineae (Caryophyllaceae) have revealed a clearer boundary between the genus Stellaria and related genera, primarily relying on the morphological characteristics of style 3, stamens 10 and petals deeply bifid. However, the newly-published species in China, which have 5 styles or ten or more lobes per petal, challenge this boundary and necessitate further studies. In this paper, we reviewed six newly-published Chinese species of Stellaria, utilising both molecular phylogenetic evidence from nuclear ribosomal internal transcribed spacer (ITS) and four plastid regions (trnL-F, matK, rbcL, rps16) and morphological evidence. Our results demonstrated that the five new species (Stellaria abaensis, S. multipartita, S. pentastyla, S. procumbens and S. zhuxiensis) were nested within the genus Stellaria, but Stellaria motuoensis was sister to the genus Schizotechium. Herein, we accepted four new Stellaria species and proposed a new combination in Schizotechium and a new synonym in Stellaria. Additionally, we described a new species Stellaria longipedicellata from Sichuan Province, China, which was distinguished from the closely-related species Stellaria decumbens by its glabrous body, linear-lanceolate leaves, long pedicellate flowers, prostrate growth habit and flowers nearly equal to or slightly shorter than sepals. Both molecular and morphological evidence support the treatment of S. longipedicellata as a new species of the genus Stellaria.
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Critical species inventories provide primary biodiversity data crucial for biogeographical, ecological, and conservation studies. After six years, a second update to the inventory of the vascular flora native to Italy is presented. It provides details on the occurrence at regional level and, for the first time, floristic data for San Marino. The checklist includes 8,241 species and subspecies, distributed in 1,111 genera and 153 families; 23 taxa are lycophytes, 108 ferns and fern allies, 30 gymnosperms, and 8,080 angiosperms. The species/subspecies endemic to Italy are 1,702, grouped in 71 families and 312 genera. The taxa currently occurring in Italy are 7,591, while 545 taxa have not been confirmed in recent times, 94 are doubtfully occurring in the country, 11 are data deficient, and 236 are reported by mistake and to be excluded at national level. Out of the 545 not confirmed taxa, 28 are considered extinct or possibly extinct.
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One of the families with the most species in the flora of Turkey, Caryophyllaceae includes about 100 genera and 3000 species worldwide. As a result of the studies, taxa classified under the subgenus Eremogone Fenzl (about 70 species) and Eremogoneastrum Williams (22 species) of Arenaria Ruppius ex L. and subgenus Spergella (Fenzl) McNeill (3 species) of Minuartia Loefl. were moved to the genus Eremogone. The genus Eremogone is perennial plants with a woody base, usually densely cushion-forming and branched. Leaves are usually clustered at or near the base of flowering stems. The genus Eremogone is distributed in north temperate regions, especially in Northwest America, Eurasian Mountains, Asia and Asia Minor with 100 taxa worldwide. In Turkey, it consists of a total of 19 taxa including 1 subspecies and 12 of them are endemic. The aim of this study was to revise the Eremogone taxa distributed in Turkey based on morphology. National and international herbarium specimens were examined by stereo microscope and living material collected through field studies were examined. Characters that can be used in the identification of taxa and species distinguishing were clarified. As a result of the investigations, it was determined that glandular hairs and the number of flowers in the inflorescence can be variable in species classification. It was determined that sepals, petals, staminal glands, the number of leaf pairs on the stem and the size of the leaves are important stable characters in the species identification keys to be used. However, especially the inward folds on the margins of the sepals cause confusion in species determination. In this study, the diagnostic characters of the taxa, the differences of the morphologically related taxa, habitat photographs and information about their distribution are given.
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Arenaria L. sensu lato is a genus of Caryophyllaceae of which around 100 species are known, distributed on all continents. The existence of four species of the genus Arenaria (Caryophyllaceae) for the vascular flora of Chile, three native and one introduced, is reported, thus reducing the historical number of names referring to this genus and accepting A. digyna Schltdl. A. rivularis Phil., A. serpens Kunth and A. serpyllifolia L., as currently accepted. New synonyms are proposed: A. microphylla Naudin, A. oligosperma Naudin, Arenaria pleurantha Phil., A. cordillerana Iamonico and A. philippii Iamonico, all of them are treated as A. serpens Kunth.
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Generic delimitation in Caryophyllaceae has been challenging and has been informed most recently by use of molecular phylogenetic data. In this study, analysis of 29 samples from the small segregate Mononeuria using nuclear ITS and plastid rps16 data revealed it to be polyphyletic. The type species, (M. patula) and two other species (M. muscorum and M. paludicola) were shown to belong to the Sabulina clade. The remaining species formed a clade that also included the previously monotypic Geocarpon and was sister to a heterogeneous group that included the Hawaiian Schiedea and three other monotypic genera, Honckenya, Wilhelmsia and Triplateia. Although several nomenclatural options are available, we propose to place the species from this clade into a single genus, Geocarpon, which basically follows the most recent treatment after exclusion of Sabulina spp., but with the necessary new genus placements. New combinations are proposed: Sabulina muscorum; Sabulina paludicola; Geocarpon carolinianum; Geocarpon cumberlandensis; Geocarpon glabrum; Geocarpon groendlandicum; Geocarpon nuttallii and Geocarpon uniflorum. Analysis of the sequence data revealed remarkable variability among populations of Sabulina patula (formerly Mononeuria patula), Sabulina paludicola (formerly Mononeuria paludicola) and Geocarpon groenlandicum (formerly Mononeuria groenlandica), suggesting that cryptic species may be present. The data also suggested that broader sampling of Sabulina and Geocarpon could lead to increased understanding of the timing and origins of occupation of calcareous glades and rock outcrop habitats in eastern North America.
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Dianthus berkayii belongs to sect. Fimbriati was described and illustrated as a new species from Erzincan Province in Eastern Anatolia, Turkey. In the current study morphological, palynological, and molecular characters were compared and discussed with its closely related species. The new species is similar to D. crinitus with fimbriate corolla, linear, acuminate, ± rigid leaves. Also similar to D. vanensis with fimbriate corolla, linear, acuminate, ± rigid, curved divaricate leaves. But distinctly different from D. crinitus by having linear and curved divaricate leaves, stem with 3–4(–5) internodes (not 5–8), a style much longer than petals, and different from D. vanensis with narrower leaves, ebarbulate and long fimbriate corolla. The bracts of D. berkayii are 1/3 of the length of the calyx, but not equal to the calyx length. The IUCN threat category of D. berkayii was determined to be “CR (Critically Endangered)” therewithal it is an Irano-Turanian element. In the study, the geographical distributions of D. berkayii, D. crinitus, and D. vanensis are also mapped, the identification key of these species was provided, pollen morphology and phylogenetic analyses based on ITS region of rDNA and ecological notes are also presented.
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The taxonomic identity and phylogenetic relationships of several South African perennial taxa often synonymised to the European Spergularia media are discussed. In particular, the case of Arenaria glandulosa Jacq., a species described as native to the Cape region of South Africa, was revisited. We found this taxon to be a Spergularia, endemic to salt-laden coastal (and rarely also saline inland) habitats of the Western Cape and Eastern Cape provinces of South Africa. This taxon is a prostrate to procumbent caespitose herb, densely glandulous all over, with small flowers solitary at the base of each leaf-like bract or in congested dichasial cymes, producing petals about equalling the sepals and free styles, among other characters. Further, we describe a morphologically similar species (often mistaken for S. media and S. bocconei) occurring in succulent Karoo scrub of Namaqualand, as a new species, named S. namaquensis. Both are glandulous dwarf subshrubs considerably differing from the recently discovered S. quartzicola, a soil-specialist endemic from Namaqualand quartzitic outcrops, by many notable vegetative and reproductive features. Molecular analyses of plastid (trnL–trnF region) and nuclear ribosomal (5.8S-ITS2 region) DNA sequence data support the morphological differentiation between the South African species S. glandulosa, S. namaquensis and S. quartzicola, representing an independent lineage when compared with the Northern Hemisphere members of the S. maritima group. Synonymy, new complete descriptions and ecological and distributional data are provided to facilitate further identification of species within the putative S. media group.
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Pseudocerastium is a monotypic genus in Caryophyllaceae endemic to China. The genus has been widely accepted since it was described in 1998, however its phylogenetic position within Caryophyllaceae has never been studied. In the present study, the whole plastid genome and nuclear ribosomal internal transcribed spacer (ITS) sequences of P. stellarioides was obtained through genome skimming, and the phylogenetic position of the species was studied for the first time. Plastid phylogenomic analysis of Caryophyllaceae revealed that Pseudocerastium is clustered within the tribe Alsineae with strong support. Phylogenetic analyses based on an enlarged taxon sampling of Alsineae using five DNA regions (matK, rbcL, rps16 intron, trnL-F and ITS) revealed that P. stellarioides was nested deeply within Cerastium with strong support. Analyses of morphological character evolution suggest that the ancestral states in Alsineae include three styles and a six-lobed capsule at the apex, while both Cerastium and Pseudocerastium have five styles and ten lobes at the apex of the capsule, further supporting their close relationship. The species P. stellarioides is similar to C. wilsonii in morphology, but differs in having villous indumentum on the lower part of the filaments and compressed globose seeds. Therefore, based on the present molecular and morphological evidence, the generic name Pseudocerastium is reduced here as a new synonym of Cerastium and the species P. stellarioides is transferred to Cerastium as C. jiuhuashanense.
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Establishing a baseline of current Arctic vascular plant diversity and distribution is critical, given the rapid and major environmental changes occurring in the Arctic ecozone in response to climate change. Here, we report the results of a floristic study of vascular plant diversity of Dorset and Mallik islands in the Canadian Arctic Archipelago, Nunavut, Canada. These two small islands lie off the coast of the Foxe Peninsula of southwestern Baffin Island, and they are part of the Circumpolar Arctic bioclimate Subzone C. The hamlet of Kinngait (formerly Cape Dorset) is located on Dorset Island, and Nunavut’s Mallikjuak Territorial Park encompasses all of Mallik Island. The study is based on a specimen-based dataset comprising 876 unique collections from the two islands gathered over the last century, including 268 new ones collected in 2015. Results are presented in an annotated checklist. The vascular plant flora of the study area comprises 26 families, 71 genera, 150 species and three infraspecific taxa; 139 species are recorded on Dorset Island and 102 on Mallik Island. Eleven taxa are newly recorded from the study area in six families: Carex rupestris, Eriophorum scheuchzeri subsp. scheuchzeri, E. triste (Cyperaceae); Diapensia lapponica (Diapensiaceae); Equisetum arvense subsp. alpestre (Equisetaceae); Oxytropis deflexa var. foliolosa (Fabaceae); Potentilla arenosa subsp. arenosa, P. hyparctica subsp. hyparctica (Rosaceae); Antennaria friesiana subsp. friesiana, Askellia pygmaea, and Taraxacum phymatocarpum (Asteraceae).
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Background: Cherleria (Caryophyllaceae) is a circumboreal genus that also occurs in the high mountains of the northern hemisphere. In this study, we focus on a clade that diversified in the European High Mountains, which was identified using nuclear ribosomal (nrDNA) sequence data in a previous study. With the nrDNA data, all but one species was monophyletic, with little sequence variation within most species. Here, we use genotyping by sequencing (GBS) data to determine whether the nrDNA data showed the full picture of the evolution in the genomes of these species. Results: The overall relationships found with the GBS data were congruent with those from the nrDNA study. Most of the species were still monophyletic and many of the same subclades were recovered, including a clade of three narrow endemic species from Greece and a clade of largely calcifuge species. The GBS data provided additional resolution within the two species with the best sampling, C. langii and C. laricifolia, with structure that was congruent with geography. In addition, the GBS data showed significant hybridization between several species, including species whose ranges did not currently overlap. Conclusions: The hybridization led us to hypothesize that lineages came in contact on the Balkan Peninsula after they diverged, even when those lineages are no longer present on the Balkan Peninsula. Hybridization may also have helped lineages expand their niches to colonize new substrates and different areas. Not only do genome-wide data provide increased phylogenetic resolution of difficult nodes, they also give evidence for a more complex evolutionary history than what can be depicted by a simple, branching phylogeny.
Chapter
This contribution provides a survey on the high-mountain vegetation of Anatolia, Turkey, covering the West Anatolian Mts., the Taurus mountain system, the Inner Anatolian volcanoes, the East Anatolian highlands and the Black Sea Mts. Due to its location between the Black Sea and the Mediterranean Sea, the intersection of three phytogeographical regions, a highly varied geologic and climatic setting in addition to a dramatic geological past, the plant life of the different Anatolian mountains systems and isolated peaks is amazingly diverse and very rich in endemics. The chapter introduces all important high-mountain ranges, their zonation, major ecosystems, key vegetation types and floristic inventories. Based on a thorough phytogeographic analysis of the high-mountain flora and an evaluation of a wide range of floristic and phylogenetic studies, the diversity patterns of the Anatolian mountain systems and their floristic links to the adjacent areas are reviewed and mapped.
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The complete plastome sequences of six species were sequenced to better understand the evolutionary relationships and mutation patterns in the chloroplast genome of the genus Colobanthus. the length of the chloroplast genome sequences of C. acicularis, C. affinis, C. lycopodioides, C. nivicola, C. pulvinatus and C. subulatus ranged from 151,050 to 151,462 bp. The quadripartite circular structure of these genome sequences has the same overall organization and gene content with 73 protein-coding genes, 30 tRNA genes, four rRNA genes and five conserved chloroplast open reading frames. A total of 153 repeat sequences were revealed. Forward repeats were dominant, whereas complementary repeats were found only in C. pulvinatus. the mononucleotide SSRs composed of A/t units were most common, and hexanucleotide SSRs were detected least often. eleven highly variable regions which could be utilized as potential markers for phylogeny reconstruction, species identification or phylogeography were identified within Colobanthus chloroplast genomes. Seventy-three protein-coding genes were used in phylogenetic analyses. Reconstructed phylogeny was consistent with the systematic position of the studied species, and the representatives of the same genus were grouped in one clade. All studied Colobanthus species formed a single group and C. lycopodioides was least similar to the remaining species.
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This is the twenty-three of a series of reports of chromosomes numbers from Mediterranean area, peri-Alpine communities and the Atlantic Islands, in English or French language. It comprises contributions on 56 taxa: Anthriscus, Bupleurum, Dichoropetalum, Eryngium, Ferula, Ferulago, Lagoecia, Oenanthe, Prangos, Scaligeria, Seseli and Torilis from Turkey by Ju. V. Shner, T. V. Alexeeva, M. G. Pimenov & E. V. Kljuykov (Nos 1768-1783); Astrantia, Bupleurum, Daucus, Dichoropetalum, Eryngium, Heracleum, Laserpitium, Melanoselinum, Oreoselinum, Pimpinella, Pteroselinum and Ridolfia from Former Jugoslavia (Slovenia), Morocco and Portugal by J. Shner & M. Pimenov (1784-1798); Arum, Biarum and Eminium from Turkey by E. Akalin, S. Demirci & E. Kaya (1799-1804); Colchicum from Turkey by G. E. Genç, N. Özhatay & E. Kaya (1805-1808); Crocus and Galanthus from Turkey by S. Yüzbaşioǧlu, S. Demirci & E. Kaya (1809-1812); Pilosella from Italy by E. Di Gristina, G. Domina & A. Geraci (1813-1814); Narcissus from Sicily by A. Troia, A. M. Orlando & R. M. Baldini (1815-1816); Allium, Cerastium, Cochicum, Fritillaria, Narcissus and Thymus from Greece, Kepfallinia by S. Samaropoulou, P. Bareka & G. Kamari (1817-1823).
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Nucleotide sequences of the plastid encoded gene matK were examined for their potential utility in phylogenetic analyses within angiosperm families. Sequences 661 bases in length were obtained from twenty species of Polemoniaceae. Phylogenetic analyses resulted in four equally parsimonious trees with a consistency index of 0.70. Several well supported groups allowed us to test hypotheses of relationship within Polemoniaceae. The segregation of Ipomopsis and Allophyllum from Gilia was supported by the placement of each in distinct groups separate from a group of four species of Gilia. Several strongly supported groups include genera now placed in different tribes. There was no support for the current separation of temperate Polemoniaceae into two tribes. The tropical genera were resolved as basal and paraphyletic within the family. The family as a whole was monophyletic with no support for the segregate family Cobaeaceae. Sequences of matK, a gene that had not been used previously for phylogenetic analyses, provided a sufficient number of reliable characters for phylogenetic analysis within Polemoniaceae. Pairwise comparisons of matK and rbcL sequences of the same taxa were performed. Sequences of matK varied at an overall rate twice that of rbcL sequences. Substitutions at the third codon position predominated in rbcL sequences, while in matK substitutions were more evenly distributed across codon positions.
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identical to print version). TAXON 62 (4) • August 2013: 831 Braun & Heuchert • (2176) Conserve Coniothecium betulinum (2176) Proposal to conserve the name Coniothecium betulinum against Torula stilbospora (Ascomycota). Lectotypus (hic designatus): without locality, on wood, without date, Corda (PRM No. 155699). Torula stilbospora Corda (in Sturm, l.c.) was later confused by Corda (Icon. Fung. 5: 50. 1842) with a fungus nowadays usually treated as "Taeniolella stilbospora (Corda) S. Hughes". Corda's (l.c. 1842: t. 2, fig. 13) second illustration of "Torula stilbospora" is quite distinct from his original drawing (Corda in Sturm, l.c.: t. 46), show-ing a fungus with a more aggregated, granular conidial structure quite distinct from Taeniolella S. Hughes which is in contrast character-ized by forming catenate conidia in firm, oblong-cylindrical chains. This confusion was already realized and discussed by Saccardo (Syll. Fung. 4: 265. 1886). Among original collections deposited in Corda's herbarium (PRM) as Torula stilbospora, there is a single collection without locality and date, but with a small pencil drawing on the envelop agreeing with Corda's original illustration, which represents true type material. This sample has been re-examined and proved to be typical Trimmatostroma betulinum (Corda) S. Hughes (≡ Coniothecium betulinum Corda), i.e., Torula stilbospora agrees with Coniothecium betulinum and is an older heterotypic synonym. Corda (l.c. 1842) cited collections from Neustadtel (locality unclear as at least four towns with this German name are known, two in the Czech Republic [now Dolny Bělá and Jezvé, part of Stružnice], one in Poland [now Nowe Miasteczko] and one in Germany [Saxony]), Senftenberg (Germany) and Prague (Czech Republic). Only the mate-rial from "Neustadtel" collected by Corda (PRM 155698) is preserved and was also re-examined. This material is very sparse, but quite different from the lectotype of Torula stilbo spora. Maintained frag-ments of conidial chains agree well with Corda's (l.c. 1842: t. 2, fig. 13) illustration and represent a genuine Taeniolella, hitherto generally known as T. stilbospora. However, the latter name is misapplied, i.e., the true Taeniolella needs another name. Trimmatostroma betulinum (Corda) S. Hughes is a common, widespread hyphomycete on bark of mostly fallen branches of Alnus, Betula, Corylus, Quercus, Pinus, Populus and Salix species mainly in Europe, but also recorded from Cuba, New Zealand and Turkey, and a widely used name in the mycological literature since its introduction by Hughes did not give any details of the original material on which he based the name C. betulinum, except for the substrate (Betula alba L.), but since only a single collection under this name is deposited in Corda's herbarium, this sample may be considered as holotype material. According to Crous & al. (in Stud. Mycol. 58: 1–32. 2007), this species belongs phylogenetically to the Dermate aceae (Helotiales, Leotiomycetes, Ascomycota). However, the older name Torula stilbospora proved to be a heterotypic synonym of Conio-thecium betulinum, the basionym of Trimmatostroma betulinum, and threatens the latter well-known name. A reallocation of Torula stilbospora, hitherto consistently used in the misapplied combination Taeniolella stilbospora, to Trimmatostroma would undoubtedly cause strong confusion. Hence, a conservation of Coniothecium betulinum over Torula stilbospora is desirable and proposed in order to maintain the current use of Trimmatostroma betulinum.
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Annual or perennial herbs, or subshrubs, rarely shrubs or small trees, usually monoecious, rarely gynodioecious or dioecious. Stems often swollen at the nodes, usually with anomalous secondary growth in older stems, also often occurring in the roots. Leaves opposite, decussate or apparently whorled, very rarely alternate, simple, entire, often connate at the base, sometimes succulent; stipules wanting, more rarely present, then mostly scarious. Flowers commonly protandrous, actinomorphic, very rarely weakly zygomorphic, usually bisexual, rarely unisexual, hypogynous or perigynous, sometimes obdiplostemonous. Inflorescences mostly dichasial, sometimes monochasial, rarely flowers solitary. Sepals (4−)5 or rarely more or fewer, imbricate, rarely valvate, completely free or connate for often most of their length, sometimes subtended by bracts (epicalyx). Petals (4−)5 or rarely more, free, aestivation contorted or rarely imbricate, entire, emarginate, bifid or lacerate, clawed or not, sometimes with coronal scales at the top of the claw, occasionally very small or absent.
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Chromosome numbers of 566 taxa (3852 individuals) of mainly South Swedish vascular plants are published, most of them previously not documented for Sweden. The plant material was collected in the late 50s by Borje Lovkvist and co-workers from natural habitats. The results were not published, although many of Lovkvist's chromosome numbers appear with an asterisk in Weimarck (1963). However, these numbers were largely based on preliminary oral information by Lovkvist and only a little more than half could be verified from Lovkvist's documented material. The entire material (field notebooks, voucher specimens, slides, laboratory journals and drawings) has been critically examined and edited by U-M. Hultgard for the Flora Nordica project. - Flora Nordica Note no. 23.
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Comparative sequencing of the maturase-encoding chloroplast gene matK has great potential for reconstructing phylogenetic relationships not only within families, but also within genera of land plants. This gene of 1550 bp is easily amplified due to highly conserved, flanking coding regions that include the trnK exons, rps 16, and psbA. Several available sequencing primers also have wide applicability. Parsimony analysis of 45 matK sequences representing Saxifragaceae sensu stricto provides a level of resolution comparable to that obtained via chloroplast DNA restriction site analysis. Furthermore, this analysis suggests relationships among genera and species that are highly concordant with the results of separate analyses of rbcL sequences and chloroplast DNA restriction sites, and with those of combined analyses of these three chloroplast DNA data sets. Parsimony analysis of 31 matK sequences representing all six sections of Gilia (Polemoniaceae) and 10 allied genera provides strong evidence for the polyphyly of Gilia and suggests relationships among sections of Gilia that are highly concordant with a recent ITS sequence analysis of the Polemoniaceae. Our analyses suggest that matK sequences are not strongly biased toward transitions, and the frequency of mutations at the first and second codon positions approach the frequency of mutations in the third codon position.
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25 populations from Turkey and one of Syria belonging to theSabulina section of the genusMinuartia have been karyologically examined. New chromosome numbers have been recorded forM. mesogitana andM. hybrida subsp.turcica, and a new variety was found in theM. hybrida complex. The origin of the taxa with n = 23 and n = 35 is discussed.
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A new apetalous species of Minuartia from the high Andes of northwestern Argentina, M. altoandina, is here described and illustrated. At first sight, M. altoandina is morphologically very similar to the European M. sedoides because of the absence of petals and ciliate leaf margin with narrow hyaline teeth; however, from a biogeographical point of view, it will probably be related to the North American M. rossii complex through the morphology of M. austromontana.
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The aim of the present study was to infer a substantially larger, more evenly sampled, phylogenetic tree for Caryophyllaceae in order to more confidently resolve relationships within this clade. This would allow us to evaluate previous classification schemes and to infer the evolution of a number of characters that have figured prominently in higher-level taxonomic treatments. We have inferred a 630-tip phylogeny (ca. 30% of the 2200 species) using maximum likelihood analyses of data from the nuclear ribosomal ITS region and five chloroplast genes and intergenic spacers: matK, ndhF, trnL-trnF, trnQ-rps16, and trnS-trnfM. Our results confirm that subfamily Paronychioideae is paraphyletic at the base of Caryophyllaceae. Alsinoideae and Caryophylloideae together form a clade, within which neither subfamily is monophyletic. With only a few exceptions, our results support the tribal classification presented by Harbaugh & al. (2010). In agreement with other recent studies, it appears that many of the larger genera are not strictly monophyletic. Our results imply that the first Caryophyllaceae had stipules, free sepals, small apetalous flowers with few stamens, and single-seeded indehiscent or irregularly dehiscing utricles. Stipules were lost along the branch to the Alsinoideae-Caryophylloideae clade, and the evolution of a tubular calyx marks Caryophylloideae. The evolution of petals, 10 stamens, and capsule fruits is inferred to have taken place along the branch subtending a clade that includes Sperguleae (mostly containing former members of Paronychioideae) and the remainder of Caryophyllaceae. As this previously unnamed major group is both well-supported in molecular phylogenetic studies and marked by clear-cut apomorphies, we propose the name Plurcaryophyllaceae for this clade and provide a phylogenetic definition.
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Minuartia dirphya from Mt Dirphys (Evvia) is described as a species new to science. It is closely related to M. wettsteinii from Mt Thriptis (East Kriti) and M. parnonia from Mt Pamon (South Peloponnisos). The three narrow endemic species form a well-defined group, the Minuartia wettsteinii-group, within Minuartia ser. Laricifoliae Mc Neill. Additions to the descriptions, chorological and ecological data for M. wettsteinii and M. parnonia are provided. The morphological characters of the group and the differences to the other species of ser. Laricifoliae are discussed. A key to the species, illustrations of the taxa and their seeds, chromosome numbers and data on their geographical distribution are also provided.
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The importance and abundance of cryptic species among invertebrate taxa is well documented. Nowadays, taxonomic, phylogenetic and conservation biological studies frequently use molecular markers to delineate cryptic taxa. Such studies, however, often face the problem of the differential resolution of the molecular markers and techniques involved. This issue is explored in the present study of cryptic taxa within the terrestrial slug complex Arion subfuscus/fuscus in continental north-west Europe. To this end, morphological, allozyme and mitochondrial 16S rDNA sequence data have been jointly evaluated. Using allozyme data and gonad type, two distinct groups were consistently delineated, even under sympatric conditions. The 16S rDNA data strongly supported both those groups and even suggested the presence of three distinct taxa within one of them. However, in view of: (1) the allopatric distribution of three OTUs, (2) the lack of allozyme or morphological differentiation, and (3) the extremely high degree of intraspecific mtDNA variation reported in pulmonate gastropods, they are, for the time being, not regarded as valid species under the biological species concept. By means of 16S rDNA and allozyme data, the position of type and topotype material of A. subfuscus s.s. and A. fuscus relative to the newly defined OTUs was determined, thus clarifying the nomenclature of this species complex. Additionally, gonad type proved to be a useful character for distinguishing the two species in north-west Europe. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 23–38.
Article
Caryophyllaceae is a principally holarctic family including around 2200 species often classified into the three subfamilies Alsinoideae, Caryophylloideae, and Paronychioideae. Complex and possibly homoplasious morphological characters within the family make taxa difficult to delimit and diagnose. To explore part of the morphological evolution within the family, we investigated the phylogeny of the Caryophyllaceae by means of analyzing plastid and nuclear sequence data with parsimony and Bayesian methods. We describe a mode of tracing a stable phylogenetic signal in ITS sequences, and a significant common signal is shared with the plastid data. Parsimony and Bayesian analyses yield some differences in tree resolution. None of the subfamilies appear monophyletic, but the monophyly of the Caryophylloideae is not contradicted. Alsinoideae are paraphyletic, with Arenaria subg. Eremogone and Minuartia subg. Spergella more closely related to the Caryophylloideae. There is strong support for the inclusion of Spergula-Spergularia in an Alsinoideae-Caryophylloideae clade. Putative synapomorphies for these groupings are twice as many stamens as number of sepals and a caryophyllad-type of embryogeny. Paronychioideae form a basal grade, where tribe Corrigioleae are sister to the rest of the family. Free styles and capsules with simple teeth are possibly plesiomorphic for the family.
Moehringia 13. Seeds exarillate 14 14. Flowers strongly perigynous
  • ........................................ Seeds Arillate.................................................................................. S And C Asia ).................................... Styles
Styles (carpels) 3–5.......................................... 22 13. Seeds arillate..................................... Moehringia 13. Seeds exarillate............................................... 14 14. Flowers strongly perigynous (S and C Asia)............................................................... Thylacospermum 14. Flowers weakly perigynous or hypogynous............. 15
19 16. Stamens 2 Arenaria 17. Capsule 2-toothed Bufonia 18. Plants minute, moss-like; flowers solitary (Chile) Reicheëlla Pax 18. Plants rush-like herbs or subshrubs; flowers not solitary (Mediterranean and Irano-Turanian regions)
  • ...................................................................................................................................................................................................................................................................................... Sepals
Sepals and petals 5........................................... 19 16. Stamens 2..................................................... 17 16. Stamens 3–8.................................................. 18 17. Capsule 4-valved or -toothed...................... Arenaria 17. Capsule 2-toothed.................................... Bufonia 18. Plants minute, moss-like; flowers solitary (Chile)........................................................... Reicheëlla Pax 18. Plants rush-like herbs or subshrubs; flowers not solitary (Mediterranean and Irano-Turanian regions).... Bufonia 19. Capsule with 2 valves or teeth................ Lepyrodiclis 19. Capsule with 4 valves or teeth............................. 20 20. Ovules and seeds numerous....................... Arenaria 20. Ovules 2(–5);
Reli-quiae haenkeanae; seu Descriptiones et icones plantarum quas in America meridionali et boreali Contributions to a chromosome atlas of the New Zealand flora – 25 miscellaneous species
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Bartling, F.G. 1831 ( " 1835 " ). Triplateia. P. 11 in: Presl, C.B., Reli-quiae haenkeanae; seu Descriptiones et icones plantarum quas in America meridionali et boreali, vol. 2(1). Prague: J.G. Calve Beuzenberg, E.J. & Hair, J.B. 1983. Contributions to a chromosome atlas of the New Zealand flora – 25 miscellaneous species. New Zealand J. Bot. 21: 13–20.
Contribution à la cytotaxono-mie du genre Minuartia L. (Caryophyllacées) en Turquie et dans quelques régions voisines
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Contribution à l’étude cytologique des genres Minuartia et Arenaria.
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Nombres chromosomique de quelque taxa principalement balkaniques du genre Minuartia (L.) Hiern
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